Filogenia das carnívoras 'tipo-orquídea' (gen. Utricularia sect. Orchidioides e Iperua: Lentibulariaceae) com observações sobre o sistema estolão-tubérculo
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Data
2017-05-19
Autores
Rodrigues, Fernanda Gomes [UNESP]
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Universidade Estadual Paulista (Unesp)
Resumo
As plantas carnívoras "tipo-orquídea" (Utricularia) compreendem 15 espécies separadas em duas seções: Orchidioides e Iperua. Essas espécies robustas e principalmente epífitas foram originalmente agrupadas dentro de sect. Orchidioides pelos primeiros sistemas taxonômicos. Estas espécies foram divididas mais tarde em duas seções quando sect. Iperua foi proposta. Devido a falta de fortes evidências apoiadas em uma perspectiva filogenética robusta, apresentamos uma proposta filogenética baseada em quatro sequências de DNA diferentes (plastidial e nuclear) e morfologia para testar a monofilia de ambas as seções. Em comparação com todos os estudos filogenéticos anteriores, cobrimos o maior número de espécies dentro das seções: 11 espécies das seções Orchidioides e Iperua com 14 espécies como grupo externo. A máxima verossimilhança e inferência Bayesiana foram aplicadas às sequências de DNA de rps16, trnL-F, matK, ITS e em três caracteres morfológicos: (1) a crista da corola, (2) os órgãos primários no embrião e (3) os tubérculos. Além disso, apresentamos uma análise histoquímica dos estolões e tubérculos sob uma perspectiva evolutiva. Nossas análises mostraram a parafília de sect. Iperua, já que Utricularia humboldtii está mais relacionada com o clado de sect. Orchidioides. Utricularia cornigera está agrupada ao clado de sect. Iperua, baseado em seqüências de cpDNA, mas é alinhada à sect. Orchidioides de acordo com o conjunto de dados ITS. Os caracteres morfológicos também não sustentam a divisão das espécies "tipo-orquídea" em duas seções. Além disso, de acordo com análises histológicas, o sistema estolão-tubérculo de ambas as seções têm como função principal o armazenamento de água, sendo esta uma importante adaptação derivada de estolões pelo menos duas vezes na história filogenética das plantas carnívoras “tipo-orquídeas”. Assim, nosso estudo fornece fortes evidências, baseadas em sequências de DNA de dois compartimentos genômicos (plastidial e nuclear) e morfologia, para agrupar as seções Utricularia sect. Orchidioides e sect. Iperua.
The “orchid-like” bladderworts (Utricularia) comprises of 15 species separated into two sections: Orchidioides and Iperua. These robust and mostly epiphytic species were originally grouped within the section Orchidioides by the first taxonomical systems. These species were later split into two sections when sect. Iperua was proposed. Due to the lack of strong evidence based on a robust phylogenetic perspective, we present a phylogenetic proposal based on four different DNA sequences (plastidial and nuclear) and morphology to test the monophyly of both sections. In comparison with all previous phylogenetic studies, we covered the largest number of species across the sections: 11 species from sections Orchidioides and Iperua with 14 species as an external group. The maximum likelihood and Bayesian inferences were applied to DNA sequences of rps16, trnL-F, matK, ITS and three morphological characters: (1) the crest of the corolla, (2) the primary organs in the embryo and (3) tubers. Additionally, we present a histochemical analysis of the stolons and tubers from an evolutionary perspective. Our analyses showed the paraphyly of sect. Iperua, since Utricularia humboldtii is more related to the clade of sect. Orchidioides. Utricularia cornigera is grouped in the sect. Iperua clade based on cpDNA sequences, but it is nested to the sect. Orchidioides according to ITS dataset. Morphological characters do not support the breaking up of the “orchid-like” species into two sections, either. Moreover, according to histological analyses, the stolon-tuber systems from both sections serves exclusively for water storage, are important adaptations and have been derived from stolons at least twice in the phylogenetic history of “orchid-like” bladderworts. Thus, our study provides strong evidence, based on DNA sequences from two genomic compartments (plastid and nucleus) and morphology to group the Utricularia sect. Orchidioides into the sect. Iperua.
The “orchid-like” bladderworts (Utricularia) comprises of 15 species separated into two sections: Orchidioides and Iperua. These robust and mostly epiphytic species were originally grouped within the section Orchidioides by the first taxonomical systems. These species were later split into two sections when sect. Iperua was proposed. Due to the lack of strong evidence based on a robust phylogenetic perspective, we present a phylogenetic proposal based on four different DNA sequences (plastidial and nuclear) and morphology to test the monophyly of both sections. In comparison with all previous phylogenetic studies, we covered the largest number of species across the sections: 11 species from sections Orchidioides and Iperua with 14 species as an external group. The maximum likelihood and Bayesian inferences were applied to DNA sequences of rps16, trnL-F, matK, ITS and three morphological characters: (1) the crest of the corolla, (2) the primary organs in the embryo and (3) tubers. Additionally, we present a histochemical analysis of the stolons and tubers from an evolutionary perspective. Our analyses showed the paraphyly of sect. Iperua, since Utricularia humboldtii is more related to the clade of sect. Orchidioides. Utricularia cornigera is grouped in the sect. Iperua clade based on cpDNA sequences, but it is nested to the sect. Orchidioides according to ITS dataset. Morphological characters do not support the breaking up of the “orchid-like” species into two sections, either. Moreover, according to histological analyses, the stolon-tuber systems from both sections serves exclusively for water storage, are important adaptations and have been derived from stolons at least twice in the phylogenetic history of “orchid-like” bladderworts. Thus, our study provides strong evidence, based on DNA sequences from two genomic compartments (plastid and nucleus) and morphology to group the Utricularia sect. Orchidioides into the sect. Iperua.
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Palavras-chave
Filogenia molecular, Morfologia, Seção Orchidioides, Seção Iperua, Tubérculo, Utricularia